Geographical parthenogenesis describes the enigmatic phenomenon that asexual organisms have larger distribution areas than their sexual relatives, especially in previously glaciated areas. Classical models suggest temporary advantages to asexuality in colonization scenarios because of uniparental reproduction and clonality. We analyzed population genetic structure and selffertility of the plant species Ranunculus kuepferi on 59 populations from the whole distribution area (European Alps, Apennines and Corsica). Amplified fragment length polymorphisms (AFLPs) and five microsatellite loci revealed individual genotypes for all populations and mostly insignificant differences between diploid sexuals and tetraploid apomicts in all measures of genetic diversity. Low frequencies of private AFLP fragments/simple sequence repeat alleles, and character incompatibility analyses suggest that facultative recombination explains best the unexpectedly high genotypic diversity of apomicts. STRUCTURE analyses using AFLPs revealed a higher number of partitions and a stronger geographical subdivision for diploids than for tetraploids, which contradicts expectations of standard gene flow models, but indicates a reduction of genetic structure in asexuals. Apomictic populations exhibited high admixture near the sexual area, but appeared rather uniform in remote areas. Bagging experiments and analyses of pollen tube growth confirmed self-fertility for pollen-dependent apomicts, but self-sterility for diploid sexuals. Facultative apomixis combines advantages of both modes of reproduction: uniparental reproduction allows for rapid colonization of remote areas, whereas facultative sexuality and polyploidy maintains genetic diversity within apomictic populations. The density dependence of outcrossing limits range expansions of sexual populations.

In temperate-zone mountains, summer frosts usually occur during unpredictable cold spells with snow-falls. Earlier studies have shown that vegetative aboveground organs of most high-mountain plants tolerate extracellular ice in the active state. However, little is known about the impact of frost on reproductive development and reproductive success. In common plant species from the European Alps (Cerastium uniflorum, Loiseleuria procumbens, Ranunculus glacialis, Rhododendron ferrugineum, Saxifraga bryoides, S. moschata, S. caesia), differing in growth form, altitudinal distribution and phenology, frost resistance of reproductive and vegetative shoots was assessed in different reproductive stages. Intact plants were exposed to simulated night frosts between −2 and −14 °C in temperature-controlled freezers. Nucleation temperatures, freezing damage and subsequent reproductive success (fruit and seed set, seed germination) were determined. During all reproductive stages, reproductive shoots were significantly less frost resistant than vegetative shoots (mean difference for LT50 −4.2 ± 2.7 K). In most species, reproductive shoots were ice tolerant before bolting and during fruiting (mean LT50 −7 and −5.7 °C), but were ice sensitive during bolting and anthesis (mean LT50 around −4 °C). Only R. glacialis remained ice tolerant during all reproductive stages. Frost injury in reproductive shoots usually led to full fruit loss. Reproductive success of frost-treated but undamaged shoots did not differ significantly from control values. Assessing the frost damage risk on the basis of summer frost frequency and frost resistance shows that, in the alpine zone, low-statured species are rarely endangered as long as they are protected by snow. The situation is different in the subnival and nival zone, where frost-sensitive reproductive shoots may become frost damaged even when covered by snow. Unprotected individuals are at high risk of suffering from frost damage, particularly at higher elevations. It appears that ice tolerance in reproductive structures is an advantage but not an absolute precondition for colonizing high altitudes with frequent frost events.

Freezing patterns in the high alpine cushion plants Saxifraga bryoides, Saxifraga caesia, Saxifraga moschata and Silene acaulis were studied by infrared thermography at three reproductive stages (bud, anthesis, fruit development). The single reproductive shoots of a cushion froze independently in all four species at every reproductive stage. Ice formation caused lethal damage to the respective inflorescence. After ice nucleation, which occurred mainly in the stalk or the base of the reproductive shoot, ice propagated throughout that entire shoot, but not into neighboring shoots. However, anatomical ice barriers within cushions were not detected. The naturally occurring temperature gradient within the cushion appeared to interrupt ice propagation thermally. Consequently, every reproductive shoot needed an autonomous ice nucleation event to initiate freezing. Ice nucleation was not only influenced by minimum temperatures but also by the duration of exposure. At moderate subzero exposure temperatures (−4.3 to −7.7 °C) the number of frozen inflorescences increased exponentially. Due to efficient supercooling, single reproductive shoots remained unfrozen down to −17.4 °C (cooling rate 6 K h−1). Hence, the observed freezing pattern may be advantageous for frost survival of individual inflorescences and reproductive success of high alpine cushion plants, when during episodic summer frosts damage can be avoided by supercooling.

SAXIFRAGA BRYOIDES L. is one of the plant species reaching the upper limits of distribution for flowering plants in the European Alps. Because of its abundance in the subnival and nival zones, we expected S. BRYOIDES to reproduce efficiently in the highly stochastic climate at higher altitudes. Investigations were carried out at two subnival sites (2650 m and 2880 m a.s.l.) in the Austrian Alps. We studied flowering phenology, dynamics of seed development, and reproductive success in the climatically different years from 2001 - 2004. For a nival plant species, S. BRYOIDES showed a particularly long prefloration period (6 - 9 weeks). From onset of anthesis until seed maturity took an individual flower another 6 - 7 weeks and all individuals at a site 9 - 10 weeks. The length of the prefloration period and seed histogenesis was temperature-dependent, whereas seed maturation seemed to be endogenously controlled. Only in the exceptionally long and warm growing season of 2003 did all fruits mature at a site. In the other years, the onset of winter conditions halted development in many fruits before maturity. The seed/ovule ratio of mature fruits was around 0.7 in all years. The relative reproductive success (RRS) ranged from zero to 0.7, depending on the site and year. In conclusion, S. BRYOIDES needs an unexpectedly long time to undergo reproductive development. Though fruit maturation is uncertain, the high S/O ratio of single intact fruits results in at least a small seed crop in most years. This seems to be sufficient to assure the spread and maintenance of S. BRYOIDES at higher altitudes. As a seed-risk strategist (Molau,1993), S. BRYOIDES would clearly benefit from a prolonged growing season, which might occur more often if climate warming continues.

Frost resistance of reproductive vs aboveground vegetative structures was determined for six common European high alpine plant species that can be exposed to frosts throughout their whole reproductive cycle. Freezing tests were carried out in the bud, anthesis and fruit stage. Stigma and style, ovary, placenta, ovule, flower stalk/peduncle and, in Ranunculus glacialis, the receptacle were separately investigated. In all species, the vegetative organs tolerated on an average 2-5 K lower freezing temperatures than the most frost-susceptible reproductive structures that differed in their frost resistance. In almost all species, stigma, style and the flower stalk/peduncle were the most frost-susceptible reproductive structures. Initial frost damage (LT₁₀) to the most susceptible reproductive structure usually occurred between -2 and -4°C independent of the reproductive stage. The median LT₅₀ across species for stigma and style ranged between -3.4 and -3.7°C and matched the mean ice nucleation temperature (-3.7 ± 1.4°C). In R. glacialis, the flower stalk was the most frost-susceptible structure (-5.4°C), and was in contrast to the other species ice-tolerant. The ovule and the placenta were usually the most frost-resistant structures. During reproductive development, frost resistance (LT₅₀) of single reproductive structures mostly showed no significant change. However, significant increases or decreases were also observed (2.1 ± 1.2 K). Reproductive tissues of nival species generally tolerated lower temperatures than species occurring in the alpine zone. The low frost resistance of reproductive structures before, during and shortly after anthesis increases the probability of frost damage and thus, may restrict successful sexual plant reproduction with increasing altitude.